Contribution to the knowledge of the genus Calcyopa Stüning, 2000 (Lepidoptera, Geometridae, Ennominae, Boarmiini), with description of a new species

#ZooKeys

ZooKeys 1233: 125-138 (2025)
DOI: 10.3897/zookeys.1233.142955

Research Article

Contribution to the knowledge of the genus Calcyopa Sttning,
2000 (Lepidoptera, Geometridae, Ennominae, Boarmiini), with
description of a new species

Bo Liu'®

1 Coconut Research Institute, Chinese Academy of Tropical Agricultural Sciences, Wenchang, Hainan 571339, China
Corresponding author: Bo Liu (liubocatas@foxmail.com)

OPEN Gaccess

Academic editor: Gunnar Brehm
Received: 28 November 2024
Accepted: 5 March 2025
Published: 28 March 2025

ZooBank: https://zoobank.
org/34D5074B-91 6B-4414-B326-
A29C4809C6C6

Citation: Liu B (2025) Contribution
to the knowledge of the genus

Calcyopa Stuining, 2000 (Lepidoptera,
Geometridae, Ennominae, Boarmiini),

with description of a new species.
Zookeys 1233: 125-138. https://doi.
org/10.3897/zookeys.1233.142955

Copyright: © Bo Liu.
This is an open access article distributed under
terms of the Creative Commons Attribution

License (Attribution 4.0 International - CC BY 4.0).

Abstract

The genus Calcyopa Stiining, 2000, is briefly reviewed. A new species, Calcyopa hainana
Liu, sp. nov., is described from Hainan Province, China. Within the genus Calcyopa, two
species groups are identified, characterized by shared traits yet distinguished by a set
of consistent features. The difoveata-group, comprises C. difoveata, C. fansipana and
C. hainana sp. nov., and the rosearia-group, includes C. rosearia, C. prasina and C. sub-
prasina. The relationship of both species groups is discussed, and an identification key
of all known Calcyopa species is presented. Illustrations are provided for adult males
and females of the difoveata-group, along with their genitalia, except for C. fansipana,
which is known only from males. DNA barcodes are provided for the type species and
the newly described species.

Key words: Calcyopa hainana, DNA barcode, identification key, new species, species
groups, taxonomic history

Introduction

The genus-group name Calichodes was first published by Warren (1897: 246)
with the description of Calichodes foveata from Penang (Peninsular Malaysia).
However, Warren did not mention it as a new genus, as the author usually did.
Therefore, Wehrli (1943: 544) considered the name to be merely a manuscript
name (“Warren M. S.”) and proposed it for a new subgenus of Boarmia, with a
different type species, B. (Calichodes) difoveata Wehrli. Subsequently, Fletcher
(1979: 32) only referenced Calichodes Wehrli, 1943, and raised it to the rank of
a genus. Holloway ([1994]: 251) redefined Calichodes Warren as a monotypic
genus, distinct from Calichodes Wehrli, and emphasized, based on a personal
communication with I. W. B. Nye, that “the earlier usage is valid under the Inter-
national Code of Zoological Nomenclature and therefore has priority”. Parsons
et al. (1999: 100) followed this treatment and mentioned only Calichodes War-
ren, 1897, but included the type species difoveata of Calichodes Wehrli, without
addressing the later invalid usage of Wehrli.

125

Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

Consequently, the generic name Calcyopa was proposed by Stiining (2000:
134) as areplacement name for Calichodes Wehrli, 1943, with Boarmia (Calicho-
des) difoveata Wehrli, 1943 designated as the type species. Additionally, Calcy-
Opa prasina Stiining, 2000 was described from Thailand, with its distribution
extending from Vietnam, Peninsular Malaysia, and Myanmar to northern India
and Nepal. Furthermore, the latter author included two other species, Ectropis
rosearia Joannis, 1929 from “Tonkin” (N. Vietnam), Yunnan (China), Thailand,
and Sumatra (Indonesia), and an undescribed species from Vietnam in this
genus. Two decades later, two new species — Calcyopa fansipana Sato, 2022
from N. Vietnam and Calcyopa subprasina Sato, 2022 from Laos — were added
to the genus. Rajaei et al. (2022) mistakenly considered Calcyopa Stiining as
a replacement name for Calichodes Warren and included all the species men-
tioned under Calichodes Warren in the catalogue of Parsons et al. (1999) under
Calcyopa Stuning, 2000.

Currently, five species of the genus Calcyopa are known, and this paper de-
scribes a new species from Hainan Island, China.

Material and methods
Specimen collection

The study is based on moth specimens housed in the following collections:
Coconut Research Institute, Chinese Academy of Tropical Agricultural Sci-
ences, Wenchang, China (CRICATAS); Institute of Zoology, Chinese Academy
of Sciences, Beijing, China (IZCAS); Natural History Museum, London, United
Kingdom (NHMUK); Institute for Agro-Environmental Sciences, NARO, Tsuku-
ba, Japan (NIAES); Zoologisches Forschungsmuseum Alexander Koenig, Bonn,
Germany (ZFMK); Zoologische Staatssammlung Minchen, Germany (ZSM).

Morphology

Terminology of the wing venation follows the Comstock-Needham System
(Comstock, 1918) as adopted for Geometridae by Scoble (1992) and Haus-
mann (2001), while genitalia terminology is based on Klots (1970) and Skou
and Sihvonen (2015). For genitalia examination, abdomens were removed and
placed in a hot 10% KOH solution. Genitalia were then dissected in 10% ethanol
and stained with Chlorazol Black E. Photographs of adult moths were taken
using a Nikon camera (D750) equipped with a Nikon lens (AF-S Micro 60 mm
f/2.8G ED). Photos of genitalia were taken with a digital camera (KUY NICE
E31SPM) attached to a Nikon microscope (SMZ745T). Focus-stacked images
were generated using Helicon Focus software (version 8.2.2 Pro).

DNA barcoding

Genomic DNA was extracted from the legs of dried adult specimens, and
the barcode fragments were amplified using primers pairs: LCO-1490 and
HCO-2198 or LepF1 and LepR1 (Folmer et al. 1994; Hebert et al. 2004). The
obtained sequences (658 bp) were deposited in the Barcode of Life Data Sys-
tems (BOLD: Ratnasingham and Hebert 2007; http://www.boldsystems.org).

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Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

All sequences mentioned in this study were obtained from the BOLD Sys-
tems. Genetic distances between species are reported as uncorrected pair-
wise distances (p-distance).

Taxonomic account

Calcyopa Stuning, 2000

Calcyopa Stiining, 2000, Moths of Nepal, Part 6. Tinea Vol. 16 (Suppl. 1):134.
Type species: Boarmia (Calichodes) difoveata Wehrli, 1943.

Generic characters. Small ennomine geometrid moths with forewing length
11-14 mm. Ground color light to dark grey or brown, with distinct black ante-
medial and postmedial lines on both wings. Head. Male antennae fasciculate,
with two pairs of shortly conical, sclerotized, ciliate projections latero-ventrally
on each segment. Cilia curved ventrad, 3-4 times longer (depending on spe-
cies) than diameter of flagellomeres. Female antennae with cilia much shorter,
the sclerotized projections absent. Vertex covered with lamellar scales. Frons
smooth-scaled, not protruding. Labial palpi slightly extending beyond frons, ba-
sally with long scales. Proboscis rather short. Chaetosemata present, near eye
margin. Thorax. Patagia and tegulae densely covered with somewhat longer
lamellar scales, distal end of tegulae with a few long hair-scales. Legs slen-
der, index of spurs 0-2-4, hind tibia not dilated, without scent brush (= hair-pen-
cil of authors) in males. Forewings moderately elongate, apex angled, termen
smoothly curved, with a double fovea in males. Antemedial and postmedial
lines conspicuous on both wings. Postmedial line on forewing curved outward
between M, and M,, with the extent of curvature varying among species; out-
side postmedial line between veins M, and CuA, or even CuA, with a dark, round,
oval or squarish patch (absent in C. rosearia only) and a faint, square pale patch
outside the strongly dentate submarginal line (absent in C. rosearia). Marginal
line consisting of small, black dots in the middle between veins, alternating with
groups of dark fringe-scales. Hindwing with apex rounded, termen minutely con-
cave between vein-ends, marginal dots rather lunulate. Tornus without distinct
spot (elongate black spot present near tornus in C. prasina and C. subprasina)
Postmedial line straighter than on forewing. Discal dot distinct, visible on both
wings, but larger on forewing. Underside with pattern paler. Venation (Fig. 1).
R, and R, coincident (distal branch of R, reduced, only R, reaching costa closer
to apex), the base of the combined veins running closely parallel to vein Sc and
the stem of R,,,. (R,) +R, and the stalk of R,, both arising from the same posi-
tion, shortly before anterior angle of cell. M, from the middle of the discoidal
vein. CuA, from shortly before posterior angle of cell. Hindwing: Sc+R, running
closely parallel to upper vein of cell for a short distance; Rs arising widely before
anterior angle of cell; M, absent, represented by a fold; CuA, from shortly before
posterior angle of cell; 3A absent. Folds through cells of both wings and those
replacing CuP in forewings and M, in hindwings very vague. Pregenital abdo-
men. Tympanal organs of moderate size, without lacinia. Sterno-tympanal pro-
cess and setal comb on third sternite absent. Lateral coremata on the posterior
portion of the third segment only present in C. difoveata and C. hainana (very
small and easily brushed off during abdominal preparation). Seventh segment

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Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

1A+2A

Figure 1. Wing venation of Calcyopa difoveata.

strongly modified, short, sternite membranous, with a pair of external corem-
atous lobes, both laterally with a dense row of fine setae; anterior invaginated
coremata, with intersegmental opening between segments 6 and 7 absent (not
“weak, scarcely invaginated” as described in StUning (2000) for C. difoveata) in
all difoveata-group species (present in rosearia-group species).

Male genitalia. Uncus deeply divided, stout, triangular, apically straight,
slightly curved ventrad in species of the difoveata-group (elongate, more deep-
ly divided, distally slender, apical part strongly bent ventrad in rosearia-group
species). Gnathos with broad, flattened lateral arms, central part strong, thorn-
shaped, elongate, slightly curved, pointed in difoveata-group species (central
part delicate, spine-shaped in rosearia-group species). Juxta small, basally a
rounded plate, extended into a narrow arm distally, slightly cup-shaped and lat-
erally bifurcate at apex. Valvae acutely triangular. Basal costal arm free, stand-
ing obliquely upright towards the upper valva margin, apically dilated, with
long bristles. Saccular process stick-like, spined at apex only (C. difoveata) or

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Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

without spines (C. hainana and C. fansipana) (spined over half or more of its
length in rosearia-group species). Aedeagus short and stout, apically with a
long, acutely triangular, sclerotized process, shaft distally with short spines or
groups of strong teeth, vesica with a small row of cornuti or a round divertic-
ulum covered with numerous spicules in difoveata-group species (aedeagus
smaller, narrower, without distal extension and without teeth on shaft and vesi-
ca without cornuti in rosearia-group species).

Female genitalia. Ovipositor short, papillae anales rounded, scarcely setose.
Needle-like sclerotization between their bases present. Anterior apophyses
short, about % length of posterior apophyses. Lamella postvaginalis a round-
ed, rather small plate in difoveata-group species (a large, sclerotized plate with
a wide rounded incision anteriorly in rosearia-group species). Introitus small,
sclerotized; posterior part of bursa sclerotized, scobinate inside; anterior part
of bursa swollen, broader than posterior part, Signum consisting of 6 to 8 chains
of small dentate projections in difoveata-group species (bursa short, rounded,
signum built of dentate projections arranged to small chains or distributed over
larger areas of the bursa surface in rosearia-group species).

Diagnosis. The genus Calcyopa currently comprises a total of six species, in-
cluding the newly described species presented in this study. These species can
be distinctly divided into two groups, mainly based on the genitalia (both male
and female) characters and the abdominal coremata. Despite their differences,
all Calcyopa species share a deeply divided uncus, similarly triangular-shaped
valvae, a free basal costal arm that is strongly setose at the apex, and a unique
stick-like saccular process with or without spines in the male genitalia. These
characters collectively set them apart from other related genera such as Par-
acalcyopa Sato, Myrioblephara Warren, and Chrysoblephara Holloway. The
closely related genus Necyopa Walker exhibits most of these traits; however,
its saccular process is replaced by a strong, longitudinal fold that extends close
to the apex of the valva. In the present study, we separate all the known Cal-
cyopa species into two groups: the C. difoveata species group, which includes
C. difoveata, C. fansipana and C. hainana; and the C. rosearia species group,
which comprises C. rosearia, C. prasina and C. subprasina. The main diagnostic
characters of these two species groups are outlined in Table 1. While the two
species groups share several common characters, they are also distinguished

Table 1. Diagnostic characters separating the Calcyopa difoveata and C. rosearia species groups.

Diagnostic characters C. difoveata species group C. rosearia species group
Intersegmental abdominal coremata between absent present
ST 6 and ST 7
Uncus moderately long, apex slightly curved rather long, apex strongly bent ventrad
ventrad
Saccular process without spines or spined at apex only spined over half its length
Aedeagus rather stout, apical sclerotization long, thin, apical sclerotization absent or
tapering; cornuti spine-like indistinct, cornuti absent
Lamella postvaginalis rather small, rounded large, with a large, round incision medio-
anteriorly
Introitus small, sclerotized large, membranous
Corpus bursa long, cylindrical, proximally slightly short, oval or squarish
inflated

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Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

by a set of stable features. For the time being, we tentatively retain all of these
species within the genus Calcyopa. However, depending on further molecular
evidence, there might be a possibility of establishing a new genus for the C.
rosearia species group or merging it with the genus Necyopa Walker.

Calcyopa difoveata (Wehrli, 1943)
Figs 2-5, 14, 15, 20

Boarmia (Calichodes) difoveata Wehrli, 1943, in: Seitz, Gross-Schmett. Erde
4 (Suppl.): 544. Type-locality: “West-Tien-Mu-Shan, Chekiang” (West Tian-
mushan, Zhejiang Province, China)

Calichodes difoveata Wehrli: Fletcher, 1979, In: Nye IWB (ed.), The Generic
Names of Moths of the World 3: 32; Parsons et al. 1999, in: Scoble, Geo-
metrid Moths of the World, A Catalogue, 1: 100.

Aethalura lushanalis Sato, 1987, Japan Heterocerists’ J. 144: 289, 290, figs 3, 4,
8. Synonymized with C. difoveata by Stiining (2000).

Calcyopa difoveata: Stiining, 2000, Tinea 16 (Suppl. 1): 135, fig. 1507; Sato
2020, in: Kishida Y, Moths of Laos, Part 1, Tinea 25 (Suppl. 2): 81, pl. 28, fig.
30; Sato 2022, Tinea 26 (3): 228, 230, figs 9, 10, 24, 36.

Type material examined. Lectotype: CHINA — Zhejiang Province ° <; Pz. Cheki-
ang, West-tien-mu-shan (= Zhejiang, West Tianmushan); 1600 m; 26 Apr. 1932;
H. Hone leg.; gen. prep. slide no. 2451-DS; ZFMK (des. Stiining, 2000).

Paralectotypes: CHINA — Zhejiang Province: 2 3, 1 9; same locality and col-
lector as lectotype; 30 Apr. 1932; ZFMK. - Guangdong Province «1 9; Lienping
(Heyuan City, Lianping County); May; H. Hone leg.; coll. Wehrli, ZFMK. Remark:
Wehrli (1943: 544) mentioned seven syntypes that were collected between 23
April and 13 June. Two female paralectotypes not included in the preceding
material examined are not in coll. ZFMK, but should be the two specimens giv-
en to coll. NHMUK in exchange (1961).

Additional material examined. CHINA — Zhejiang Province « 4 <3’, 2 29; same
locality and collector as lectotype; 3 Apr., 4 Apr., 10 Apr., 17 Apr. (2), 18 Apr. 1932;
ZFMK * 5 4, 2 29; same locality and collector as lectotype; 400 m; mid-April
1936; gen. glycerol no. 6/B1<4; ZFMK « 1 9; Chekiang, Wenchow (= Zhejiang, Wen-
zhou); 18 Apr. 1939; H. Hone leg.; ZFMK « 3 34, 1 9; same locality and collector as
for preceding; June 1939; ZFMK + 1 4,2 29; same locality and collector as for pre-
ceding; July 1939; gen. glycerol no. 6/B39; ZFMK. — Fujian Province «1 <; Fukien,
Kuatun (= Fujian, Guadun); 27.40°N, 117.40°E; 2300 m; 3 Apr. 1938; J. Klapperich
leg.; ZFMK. — Guangdong Province « 2 3, 3 29; Canton or Lienping; ZFMK. -
Taiwan Province : 2 <3 (paratypes of Aethalura lushanalis Sato, 1987); Nantou
Hsien, Lushan-wenchuan; 13-14 Aug. 1983; R. Sato leg.; ZFMK * 2 9° 9; Kaohsiung,
Liukuei Sanping; 650 m; 24-26 Jul. 1987; R. Sato leg.; ZFMK * 1 3; Nantou Co.,
Hueisun Forest; 600 m; 22 Jun. 1993; F. Aulombard & J. Plante leg.; ZFMK * 1 3,
1 9; same locality and collectors as for preceding; 570-800 m; 28/29 Sep. 1992;
ZFMK. — Hainan Province «1 3, 1 2; Wuzhishan; 1333 m; 10 Jan. 2008; V. Siniaev
leg.; ZSM +5 34,3 99; Wuzhishan; 756 m; 25 Mar. 2023; Bo Liu leg.; CATASCRI.

Diagnosis. The diagnostic characters are given under the newly described
species.

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Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

side 4 female, upperside 5 ditto, underside 6 C. fansipana, holotype, male, upperside, N. Vietnam, NIAES 7 ditto, underside
8-11 C. hainana sp. nov., Hainan, China, CRICATAS/ IZCAS 8 holotype, male, upperside 9 ditto, underside 10 paratype,
female, upperside 11 ditto, underside 12, 13 specimens of C. hainana sp. nov. in resting position, type series, Hainan,
China 12 male 13 female. Scale bar: 1 cm.

Distribution. China (Zhejiang, Fujian, Jiangxi, Guangdong, Hainan, Taiwan),
Vietnam, Laos.

Genetic data. The Barcode Index Number for Calcyopa difoveata is BOLD:
AAH0723 (N = 3, Sample IDs: CRICATAS00104, BC ZSM Lep 16015, BC ZSM
Lep 15995).

Remarks. This species is now recorded for the first time from Hainan Island,
China. Some specimens from Hainan show minor differences in genitalia com-
pared to those from Taiwan and Vietnam, including variation in the thickness of
the gnathos in male genitalia as well as the shape of the lamella postvaginalis
in female genitalia. At present, we treat these as infrasubspecific variations.

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Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

Calcyopa fansipana Sato, 2022
FIGS OF7;, iG IZ

Calcyopa fansipana Sato, 2022, Tinea 26 (3): 226, figs 1, 2, 23. Type-locality: Mt
Fan-si-pan, N. Vietnam.

Type material examined. Holotype: VIETNAM * 3; N. Vietnam, Cha-pa (Sa Pa),
Mt Fan-si-pan, N. Seite; 22°17'N, 103°44'E; 1600 m; 21-23 Apr. 1995; leg. Sin-
jaev & Sammler, ex coll. A. SchintImeister; Prim Urwald; NIAES.

Paratype: VIETNAM * 1 3; same data as holotype; NIAES.

Additional material examined. VIETNAM ° 5 4; same locality and elevation
as holotype; 01-05 Mar. 1995; ex coll./ leg. Dr. R. Brechlin; ZFMK * 1 4; same
locality and collector as for preceding; 1520 m; ZFMK.

Diagnosis. Diagnostic characters are given under the newly described spe-
cies. The male adult and genitalia are illustrated in Sato (2022), but we depict
them here again for comparison with the other two species of difoveata-group.
The female is still unknown at present.

Distribution. Vietnam.

Genetic data. No data available.

Remarks. Stiining (2000: 135) already mentioned this species as “an unde-
scribed species also from Vietnam’, which was later described as fansipana
by Sato (2022: 226). He wrote in the description: “Third abdominal sternite
with setal comb” and “Hind tibia with hair-pencil”. These two characters were
mentioned by an unexplainable error (Sato 2023, pers. comm.). Both setal
comb and hair-pencil are absent in all species of the genus Calcyopa, which, of
course, was known to Sato.

Calcyopa hainana sp. nov.
https://zoobank.org/8AC4C54D-B7E3-4F34-B81 1-E5AB81F959CF
Figs 8-13, 18, 19, 21

Type material. Holotype: CHINA — Hainan Province « <'; Lingshui, Diaoluoshan;
922 m; 16-19 Apr. 2024; Bo Liu & Wei Yan leg.; CRICATAS/ IZCAS.

Paratypes: (12 3, 21 29) CHINA — Hainan Province + 1 3; Wuzhishan;
1333 m; V. Siniaev leg.; 10 Jan. 2008; ZSM * 2 3d, 2 29; same locality as
holotype; 20 Apr. 2023; Bo Liu leg.; gen. prep. nos. CRICATASO0101, CRICA-
TAS00103; CRICATAS/ IZCAS * 4 299; same locality and collectors as holotype;
05-07 Mar. 2024; CRICATAS/ IZCAS + 1 3, 1 2; same locality as holotype; 01-
03 Apr. 2024; Bo Liu, Wei Lin & Miaofeng Xu leg.; CRICATAS/ IZCAS : 4 3.3, 5
2; same data as holotype; CRICATAS/ IZCAS : 4 33,9 29; same locality and
collectors as holotype; 07-12 May 2024; CRICATAS/ IZCAS/ ZFMK.

Diagnosis. This species closely resembles C. difoveata and, though much
less, also C. fansipana, but can be distinguished by several morphological char-
acters: The postmedial line of the forewing between veins M, and M, exhibits a
more pronounced outward curve than in C. difoveata, similar to that in C. fansi-
pana. Moreover, the light-brown ground color of the upperside and the absence
of broad, dark marginal bands on the underside of the latter are also diagnostic.
The free costal arm of the valva is longer and narrower in C. hainana than in

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Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

19
Figures 14-19. Male genitalia of Calcyopa species. 14, 15 C. difoveata, Hainan, China 14 genitalia capsule 15 aedeagus,
close-up of cornuti at the bottom left 16, 17 C. fansipana, holotype, N. Vietnam, gen. prep. no. RS8731 16 genitalia cap-
sule 17 aedeagus 18, 19 C. hainana sp. nov., paratype, Hainan, China, gen. prep. no. CRICATASO0101 18 genitalia capsule
19 aedeagus. Scale bar: 1 mm.

the other two congeners and the saccular process lacks spines. The triangular
apex of the aedeagus is more pointed and elongated compared to C. difove-
ata and C. fansipana. Additionally, the cornuti in C. hainana are composed of

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Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

a group of long juxtaposed spines, while in C. difoveata they consist of shorter
spines, and in C. fansipana they form a dense cluster of minute spines covering
a round diverticulum of the vesica. A row of rather large spines and a big single
tooth present apically on the shaft of aedeagus of C. difoveata are replaced by
a large number of smaller denticles in C. hainana.

Description. Forewing length 11.1-11.9 mm in males, 11.7-14.3 mm in fe-
males. Ground color brownish-green in fresh specimens, but gradually fading
to brown over time, with numerous scattered dark scales and distinct dark an-
temedial and postmedial lines on both wings. Head. Male antennae fasciculate,
with two pairs of shortly conical, sclerotized, ciliate projections latero-ventrally
on each segment. Female antennae with cilia much shorter, and sclerotized
projections absent. The longest cilia are about 3 times the diameter of the fla-
gellum segments in males and about 0.5 times the diameter of the flagellum
segments in females. Vertex covered with lamellar, pale brown scales. Frons
smooth-scaled, not protruding. Labial palpi slightly extending beyond frons,
basally with long, dark scales, third segment small. Proboscis rather short.
Chaetosemata present, near eye margin. Thorax. Patagia and tegulae densely
covered with somewhat longer lamellar scales, distal end of tegulae with a few
long hair-scales. Wings brownish, scattered with numerous dark scales. Fore-
wings moderately elongate, apex angled, termen smoothly curved, with a dou-
ble fovea in males. Antemedial line fine, visible. Discal spot small, streak-like.
Postmedial line outcurved between veins M, and M,, at CuA,, and at 1A+2A.
Outside postmedial line between M, and CuA, with a round black patch ante-
riorly and a square pale patch posteriorly. Subterminal line zigzag, white, with
a dark inner edge. Terminal line black, with a series of dark spots between the
veins. Hindwings with apex rounded, termen minutely concave between vein-
ends. Basal area densely covered with numerous black scales. Antemedial line
broad, outcurved at lower cell vein and inner margin. Discal spot small, streak-
like. Postmedial line outcurved between veins M, and M,, and at inner margin.
Subterminal zigzag, white, with a dark edge inside. Terminal line black, with
dark spots between the veins. Underside blurry and paler, with a broad, dark
band outside postmedial line on both wings. Antemedial line faint on forewing,
but clearly visible on hindwing. Postmedial line barely outcurved on both wings.
Legs slender, index of spurs 0-2-4, hind tibia not dilated, without scent brush in
males. Pregenital abdomen. Tympanal organs, sterno-tympanal process, setal
comb, and abdominal coremata as mentioned in the generic description. Abdo-
men laterally with several pairs of scale brushes on segments 2, 3, 4, 5, 6 and 7.

Male genitalia. Uncus deeply divided, triangular, apical part straight,
slightly curved ventrad. Gnathos with broad, flattened lateral arms, central
part strong, thorn-shaped, elongate, slightly curved. Valvae acutely triangu-
lar. Basal costal arm free, standing obliquely upright towards the upper valva
margin, apically dilated, with long bristles. Saccular process stick-like, with-
out spines. Juxta small, basally a rounded plate, extended into a narrow arm
distally, slightly cup-shaped and laterally bifurcate at apex. Aedeagus short
and stout, apically with a long, acutely triangular, sclerotized process; shaft
distally with a group of small serrate projections; vesica with a small row of
long-spined cornuti. Bulbus ejaculatorius long, about four times as long as
the aedeagus shaft.

ZooKeys 1233: 125-138 (2025), DOI: 10.3897/zookeys.1233.142955 134

Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

Figures 20, 21. Female genitalia of Calcyopa species. 20 C. difoveata, Hainan, China 21 C.
hainana sp. nov., paratype, Hainan, China, gen. prep. no. CRICATASO0103. Scale bar: 1 mm.

Female genitalia. Ovipositor short, papillae anales scarcely setose. Anterior
apophyses short, about % length of posterior apophyses. Ventral longitudinal
sclerotization needle-like, basally dilated, slightly triangular. Lamella postvag-
inalis small, round or oval, bottom concave at center. Introitus bursae small,
sclerotized, significantly narrower than the posterior part of bursa, setting into
a small sternite pocket. Posterior part of bursa slightly longer than anterior
part, posterior one-third sclerotized, scobinate inside. Anterior part of bursa
swollen, triangular, noticeably broader than posterior part, consisting of 7 to 8
chains of small serrate projections on both opposed inner sides.

Etymology. The specific name of “hainana” is derived from the type locality,
Hainan Island, China.

Distribution. China (Hainan).

Genetic data. The Barcode Index Number for Calcyopa hainana is BOLD:
AAH2362 (N = 2, Sample IDs: CRICATASO0101, BC ZSM Lep 16053). The genetic
distance of C. hainana from C. difoveata (N = 3, Sample IDs: CRICATASO0104, BC
ZSM Lep 16015, BC ZSM Lep 15995) ranges from 5.32% to 5.62% (p-distance).

ZooKeys 1233: 125-138 (2025), DOI: 10.3897/zookeys.1233.142955 135

Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

Key to all known Calcyopa species, based on male and female genitalia

1

Uncus stout, deeply bifid until half its length, moderately long, apically
slightly curved ventrad; saccular process without spines or spined at apex
only; aedeagus large, rather stout, apical sclerotization long, tapering; cor-
nuti spine-like; corpus bursae long, cylindrical, proximally slightly inflated;
lamella postvaginalis rather small, rounded................. 2 (difoveata-group)
Uncus rather long and slender, deeply bifid until % of its length, apically
strongly bent ventrad; saccular process spined over half its length; aedea-
gus small, finer, apical sclerotization indistinct; cornuti absent; corpus bur-
sae short, inflated, triangular or squarish; lamella postvaginalis large, rect-
angular, with a large, round incision medio-anteriorly ...... 4 (rosearia-group)
Saccular process spined at the tip................eeeeseeeeees Calcyopa difoveata
Saccular process less developed, without SPINES .............cccceceeseeeeeeeeeees 3
Costal arm of valva slender; cornuti a group of long, juxtaposed spines...
Restore citode Dtutaniea sea peninetauss Tekxens mee Dea Atess wes aupciiNnEt nies cased ee ats C. hainana sp. nov.
Costal arm of valva stouter; cornuti smaller, arranged in two groups.........
Fe AE AVENE «DR At hia RR Oem RIEL OVERESTIMATES C. fansipana
Signum a small group of dentate processes, partly arranged in short
DIOS re bien en se AAG eanikany ATENEO assem RRh Ree mseunad 1 REY Monee iat C. rosearia
Signum consisting of two groups of spines or oblique grooves................ 5
Signum consisting of two groups of many minute spines, covering large
GkeAaS OT OURS anes A Beet eee Bsc teses 52d ee ti Po os ah meter tos C. subprasina
Signum consisting of two oblique grooves covered with spines internally...
taunted satan Renna een ry avs re onmene ate Me sanyee NN api ial ng dds age TRARNRC ORE Eva acened C. prasina

Acknowledgements

| would like to express my sincere thanks to Dieter Stiining (ZFMK, Germany) who
sent me images and data of the original specimens used by Eugen Wehrli for the
description of “Calichodes” difoveata (including type-material), to Rikio Sato (Ni-
igata, Japan) for providing me with valuable literature and images of the holotype
of Calcyopa fansipana, and to Axel Hausmann (ZSM, Germany) who sent me the
data of the Calcyopa specimens. | would also like to thank Wei Yan (CRICATAS,
China) and Jiexiong Fu (Diaoluoshan Division of Hainan Tropical Rainforest Na-
tional Park Bureau, Lingshui, China) for their assistance in specimen collection.

Additional information

Conflict of interest

The author has declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by the Science and Technology Special Fund of Hainan Province
(ZDYF2022XDNY214) and the Hainan Tropical Rainforest Conservation Research Project,
ZDYF2023RDYLO1 (supported by the Hainan Institute of National Park, HINP KY-24ZK02).

ZooKeys 1233: 125-138 (2025), DOI: 10.3897/zookeys.1233.142955 136

Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

Author contributions
Writing - Original draft: BL. Writing - Review and Editing: BL.

Author ORCID
Bo Liu © https://orcid.org/0009-0008-7003-4659

Data availability

All of the data that support the findings of this study are available in the main text.

References

Comstock JH (1918) The Wings of Insects. Comstock Publishing Company, Ithaca, 430 pp.

Fletcher DS (1979) Geometroidea. In: Nye IWB (Ed.) The Generic Names of Moths of
the World (Vol. 3). British Museum (Natural history), London, 243 pp. https://doi.
org/10.5962/bhI.title.119424

Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of
mitochondrial cytochrome c oxidase subunit | from diverse metazoan invertebrates.
Molecular Marine Biology and Biotechnology 3(5): 294-299.

Hausmann A (2001) Introduction. Archiearinae, Orthostixinae, Desmobathrinae, Also-
philinae, Geometrinae. In: Hausmann A (Ed.) The Geometrid Moths of Europe (Vol.
1). Apollo Books, Stenstrup, 282 pp.

Hebert PDN, Penton EH, Burns JM, Janzen DH, Hallwachs W (2004) Ten species in one:
DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes
fulgurator. Proceedings of the National Academy of Sciences of the United States of
America 101: 14812-14817. https://doi.org/10.1073/pnas.0406166101

Holloway JD (1994) The Moths of Borneo, Part 11: Family Geometridae, subfamily En-
nominae. Malayan Nature Journal 47: 1-309. [pls 1-19, 593 figs]

Joannis de J (1929) Lépidoptéres Héterocéres du Tonkin. Annales de la Societe Ento-
mologique de France 98: 361-552, 559-560. https://doi.org/10.1080/21686351.19
29.12280113

Klots AB (1970) Lepidoptera. In: Tuxen SL (Ed.) Taxonomists’ glossary of genitalia in
insects. Munksgaard, Copenhagen, 115-130.

Parsons MS, Scoble MJ, Honey MR, Pitkin LM, Pitkin BR (1999) The catalogue. In: Scoble
MJ (Ed.) Geometrid Moths of the World: A Catalogue (Lepidoptera, Geometridae) (Vols
1-2). CSIRO, Collingwood, 1016 pp. https://doi.org/10.1163/9789004542006_006

Rajaei H, Hausmann A, Scoble M, Wanke D, Plotkin D, Brehm G, Murillo-Ramos L, Sihvonen
P (2022) An online taxonomic facility of Geometridae (Lepidoptera), with an overview
of global species richness and systematics. Integrative Systematics: Stuttgart Con-
tributions to Natural History 5(2): 145-192. https://doi.org/10.18476/2022.577933

Ratnasingham S, Hebert PDN (2007) BOLD: The Barcode of Life Data System (http://
www.barcodinglife.org). Molecular Ecology Notes 7(3): 355-364. https://doi.
org/10.1111/j.1471-8286.2007.01678.x

Sato R (1987) Two new species of the Ennominae (Geometridae) from Taiwan. Japan
Heterocerists’ Journal 144: 289-292.

Sato R (2020) Geometridae (Ennominae, Boarmiini). In: Kishida Y (Ed.) Moths of Laos,
Part 1, Tinea 25(Suppl. 2): 60-86. [pls 18-29]

Sato R (2022) Descriptions of two new species of the genus Calcyopa Stiining and its
related new genus from Southeast Asia (Geometridae, Ennominae). Tinea 26(3):
226-234.

ZooKeys 1233: 125-138 (2025), DOI: 10.3897/zookeys.1233.142955 137

Bo Liu: Contribution to the knowledge of Calcyopa, with description of a new species

Scoble MJ (1992) The Lepidoptera: Form, Function and Diversity. Oxford University
Press, Oxford, 404 pp.

Skou P, Sihvonen P (2015) Ennominae I. In: Hausmann A (Ed.) The Geometrid Moths of
Europe (Vol. 5). Brill, Leiden, 657 pp. https://doi.org/10.1163/9789004265738

Stiining D (2000) Additional notes on the Ennominae of Nepal, with descriptions of eight
new species (Geometridae). In: Haruta T (Ed.) Moths of Nepal, part 6, Tinea 16(Suppl.
1): 94-152. [pls 170-172]

Warren W (1897) New genera and species of Drepanulidae, Thyrididae, Epiplemidae,
Uraniidae, and Geometridae in the Tring Museum. Novitates Zoologicae 4: 195-2672.

Wehrli E (1939-1954) Subfamilie: Geometrinae. In: Seitz A (Ed.) Die Grossschmetterlinge
der Erde. Vol. 4 (Supplement), Alfred Kernen Verlag, Stuttgart, 254-766. [pls 19-53]

ZooKeys 1233: 125-138 (2025), DOI: 10.3897/zookeys.1233.142955 138